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Woody encroachment is a widespread and acute phenomenon affecting grasslands and savannas worldwide. We performed a meta-analysis of 29 studies from 13 different grassland/savanna communities in North America to determine the consequences of woody encroachment on plant species richness. In all 13 communities, species richness declined with woody plant encroachment (average decline = 45%). Species richness declined more in communities with higher precipitation ( r 2 = 0.81) and where encroachment was associated with a greater change in annual net primary productivity (ANPP; r 2 = 0.69). Based on the strong positive correlation between precipitation and ANPP following encroachment ( r 2 = 0.87), we hypothesize that these relationships occur because water-limited woody plants experience a greater physiological and demographic release as precipitation increases. The observed relationship between species richness and ANPP provides support for the theoretical expectation that a trade-off occurs between richness and productivity in herbaceous communities. We conclude that woody plant encroachment leads to significant declines in species richness in North American grassland/savanna communities.

Aim: Biomes worldwide are shifting with global change. Biomes whose extents are limited by temperature or precipitation, such as the tundra and savanna, may be particularly strongly affected by climate change. While woody plant encroachment is prevalent across both biomes, its relationship to temperature and precipitation change remains unknown. Here, we quantify the degree to which woody encroachment is related to climate change and identify its main associated drivers. Location: Tundra and savanna biomes. Time period: 1992 ± 20.27–2010 ± 5.62 (mean ± SD). 1876–2016 (range). Major taxa studied: Woody plants (shrubs and trees). Methods: We compiled a dataset comprising 1,089 records from 899 sites of woody plant cover over time and attributed drivers of woody cover change across these two biomes. We calculated cover change in each biome and assessed the degree to which cover change corresponds to concurrent temperature and precipitation changes using multiple climate metrics. Finally, we conducted a quantitative literature review of the relative importance of attributed drivers of woody cover change. Results: Woody encroachment was widespread geographically and across climate gradients. Rates of woody cover change (positive or negative) were 1.8 times lower in the tundra than in the savanna (1.8 vs. 3.2%), while rates of woody cover increase (i.e., encroachment) were c. 1.7 times lower in the tundra compared with the savanna (3.7 vs. 6.3% per decade). In the tundra, magnitudes of woody cover change did not correspond to climate, while in the savanna, greater cover change corresponded with increases in precipitation. We found higher rates of woody cover change in wetter versus drier sites with warming in the tundra biome, and higher rates of woody cover change in drier versus wetter sites with increasing precipitation in the savanna. However, faster rates of woody cover change were not associated with more rapid rates of climate change across sites, except for maximum precipitation in the savanna. Main conclusions: Woody encroachment was positively related to warming in the tundra and increased rainfall in the savanna. However, cover change rates were not predicted by rates of climate change, which can be partially explained by climate interactions in both biomes. Additional likely influences include site-level factors, time-lags, plant-specific responses, and land use and other non-climate drivers. Our findings highlight the complex nature of climate change impacts in biomes limited by seasonality, which should be accounted for to realistically estimate future responses across open biomes under global change scenarios.

The book describes an integrated theory that links estuary shape to tidal hydraulics, tidal mixing and salt intrusion.The shape of an alluvial estuary is characterised by exponentially varying width and the absence of bottom slope.

Ecosystems with alternative attractors are susceptible to abrupt regime shifts that are often difficult to predict and reverse. In this study, we quantify multiple system dynamics to determine whether the transition of mesic grassland to shrubland, a widespread phenomenon, represents a linear reversible process, a nonlinear but reversible threshold process, or a transition between alternative attractors that is nonlinear and prone to hysteresis. Using a 28-yr data set with annual resolution and extensive spatial replication, we found that shrub cover is correlated with distinct thresholds of fire and C 4 grass cover, resulting in temporal bimodality of shrub cover and abrupt shifts of shrub cover despite gradual changes in grass cover. These abrupt increases in shrub cover are the most rapid ever reported in grasslands, and illustrate internal thresholds that separate grasslands and shrublands. Nonlinear transitions from low to high shrub cover were also closely associated with positive feedback mechanisms that alter fire and competition ( r 2 = 0.65), suggesting that grasslands and shrublands could show hysteresis, and by definition exist as alternative attractors. Thus, the response of this ecosystem to anthropogenic activity should tend to be rapid, nonlinear, and perhaps difficult to reverse. Regime shifts in this mesic grassland were predictable: we found that grassland and shrubland attractors were differentiated by critical thresholds of ∼50-70% grass cover, 5-10% shrub cover, and a fire return interval of ∼3 yr. These thresholds may provide adaptive potential for managing nonlinear behavior in socio-ecological systems in a changing environment.

Woody-plant encroachment is a global phenomenon that has been affecting the southwestern United States since the late 1800s. Drought, overgrazing, herbivory, and competition between grasses and shrub seedlings have been hypothesized as the main drivers of shrub establishment. However, there is limited knowledge about the interactions among these drivers. Using a rainfall manipulation system and various herbivore exclosures, we tested hypotheses about how precipitation (PPT), competition between grasses and shrub seedlings, and predation affect the germination and first-year survival of mesquite (Prosopis glandulosa), a shrub that has encroached in Southern Great Plains and Chihuahuan Desert grasslands. We found that mesquite germination and survival (1) increased with increasing PPT, then saturated at about the mean growing season PPT level, (2) that competition between grasses and shrub seedlings had no effect on either germination or survival, and (3) that herbivory by small mammals decreased seedling establishment and survival, while ant granivory showed no effect. In addition to its direct positive effect on survival, PPT had an indirect negative effect via increasing small mammal activity. Current models predict a decrease in PPT in the southwestern United States with increased frequency of extreme events. The non-linear nature of PPT effects on Mesquite recruitment suggests asymmetric responses, wherein drought has a relatively greater negative effect than the positive effect of wet years. Indirect effects of PPT, through its effects on small mammal abundance, highlight the importance of accounting for interactions between biotic and abiotic drivers of shrub encroachment. This study provides quantitative basis for developing tools that can inform effective shrub management strategies in grasslands and savannas.

Aim: Across the planet, grass-dominated biomes are experiencing shrub encroachment driven by atmospheric CO2 enrichment and land-use change. By altering resource structure and availability, shrub encroachment may have important impacts on vertebrate communities. We sought to determine the magnitude and variability of these effects across climatic gradients, continents, and taxa, and to learn whether shrub thinning restores the structure of vertebrate communities. Location: Worldwide. Time period: Contemporary. Major taxa studied: Terrestrial vertebrates. Methods: We estimated relationships between percentage shrub cover and the structure of terrestrial vertebrate communities (species richness, Shannon diversity and community abundance) in experimentally thinned and unmanipulated shrub-encroached grass-dominated biomes using systematic review and meta-analyses of 43 studies published from 1978 to 2016. We modelled the effects of continent, biome, mean annual precipitation, net primary productivity and the normalized difference vegetation index (NDVI) on the relationship between shrub cover and vertebrate community structure. Results: Species richness, Shannon diversity and total abundance had no consistent relationship with shrub encroachment and experimental thinning did not reverse encroachment effects on vertebrate communities. However, some effects of shrub encroachment on vertebrate communities differed with net primary productivity, amongst vertebrate groups, and across continents. Encroachment had negative effects on vertebrate diversity at low net primary productivity. Mammalian and herpetofaunal diversity decreased with shrub encroachment. Shrub encroachment also had negative effects on species richness and total abundance in Africa but positive effects in North America. Main conclusions: Biodiversity conservation and mitigation efforts responding to shrub encroachment should focus on low-productivity locations, on mammals and herpetofauna, and in Africa. However, targeted research in neglected regions such as central Asia and India will be needed to fill important gaps in our knowledge of shrub encroachment effects on vertebrates. Additionally, our findings provide an impetus for determining the mechanisms associated with changes in vertebrate diversity and abundance in shrub-encroached grass-dominated biomes.

The vegetation of semi‐arid and arid landscapes is often comprised of mixtures of herbaceous and woody vegetation. Since the early 1900s, shifts from herbaceous to woody plant dominance, termed woody plant encroachment and widely regarded as a state change, have occurred world‐wide. This shift presents challenges to the conservation of grassland and savanna ecosystems and to animal production in commercial ranching systems and pastoral societies. Dryland management focused on cattle and sheep grazing has historically attempted to reduce the abundance of encroaching woody vegetation (hereafter, ‘brush management’) with the intent of reversing declines in forage production, stream flow or groundwater recharge. Here, we assess the known and potential consequences of brush management actions, both positive and negative, on a broader suite of ecosystem services, the scientific challenges to quantifying these services and the trade‐offs among them. Our synthesis suggests that despite considerable investments accompanying the application of brush management practices, the recovery of key ecosystem services may be short‐lived or absent. However, in the absence of such interventions, those and other ecosystem services may be compromised, and the persistence of grassland and savanna ecosystem types and their endemic plants and animals threatened. Addressing the challenges posed by woody plant encroachment will require integrated management systems using diverse theoretical principles to design the type, timing and spatial arrangement of initial management actions and follow‐up treatments. These management activities will need to balance cultural traditions and preferences, socio‐economic constraints and potentially competing land‐use objectives. Synthesis. Our ability to predict ecosystem responses to management aimed at recovering ecosystem services where grasslands and savannas have been invaded by native or exotic woody plants is limited for many attributes (e.g. primary production, land surface–atmosphere interactions, biodiversity conservation) and inconsistent for others (e.g. forage production, herbaceous diversity, water quality/quantity, soil erosion, carbon sequestration). The ecological community is challenged with generating robust information about the response of ecosystem services and their interactions if we are to position land managers and policymakers to make objective, science‐based decisions regarding the many trade‐offs and competing objectives for the conservation and dynamic management of grasslands and savannas.

- We used a functional trait-based approach to assess the impacts of aridity and shrub encroachment on the functional structure of Mediterranean dryland communities (functional diversity (FD) and community-weighted mean trait values (CWM)), and to evaluate how these functional attributes ultimately affect multifunctionality (i.e. the provision of several ecosystem functions simultaneously).- Shrub encroachment (the increase in the abundance/cover of shrubs) is a major land cover change that is taking place in grasslands worldwide. Studies conducted on drylands have reported positive or negative impacts of shrub encroachment depending on the functions and the traits of the sprouting or nonsprouting shrub species considered. - FD and CWM were equally important as drivers of multifunctionality responses to both aridity and shrub encroachment. Size traits (e.g. vegetative height or lateral spread) and leaf traits (e.g. specific leaf area and leaf dry matter content) captured the effect of shrub encroachment on multifunctionality with a relative high accuracy (r2 = 0.63). FD also improved the resistance of multifunctionality along the aridity gradient studied.- Maintaining and enhancing FD in plant communities may help to buffer negative effects of ongoing global environmental change on dryland multifunctionality.